The Fishes

of the

Central Canadian Rockies Ecosystem

David W. Mayhood

Report No. 950408 April 1995

FWRFreshwater Research Limited

The Fishes

of the

Central Canadian Rockies Ecosystem

David W. Mayhood

Prepared forBanff National ParkBox 900Banff, Alberta T0L 0C0

by

FWR

Freshwater Research Limited

1715 Seventh Avenue NW Calgary, Alberta, Canada T2N 0Z5 Phone (403) 283-8865 Fax: (403) 283-9446

Freshwater Research Limited Report No. 950408 April 1995

© 1995 David W. MayhoodAll rights reserved

This report is the intellectual property of the author. It has not been peer-reviewed and does notconstitute a primary publication. The author may eventually publish parts of the report in similar ordifferent form in the primary literature. Unpublished material cited herein is the property of theperson, company or agency cited, who retain all rights to it, including prior right of publication.

This report may be copied freely, provided that the copies are complete and identical photocopies,specifically including the cover, title page, this copyright notice, acknowledgements and anyappendices. This report may be cited freely. Excerpts may be used, but must be clearly attributedto this document. Those wishing to publish excerpts that include the unpublished material of theauthor or others cited herein must obtain the permission of the owners of that material.

suggested citation format

Mayhood, D. W. 1995. The fishes of the Central Canadian Rockies Ecosystem. FreshwaterResearch Limited Report No. 950408 prepared for Parks Canada, Banff National Park, P. O.Box 900, Banff, AB T0L 0C0. 59 p.

Abstract

I surveyed existing information in historical documents; technical publications andreports; selected file data held by Parks Canada, Alberta Fish and Wildlife and the BritishColumbia Ministry of Environment Lands and Parks; and interviewed reliable observersknowledgeable about selected waters to determine the native and present distribution offishes within the Central Canadian Rockies Ecosystem (CCRE).

Fifty-four species of fishes in 12 families have been reported to exist in or near the CCRE.Of these, only 33 are, or once were, native to some part of the study area or in connectedadjacent waters. At least 10 native fishes have suffered impacts of significantconservation concern. One native subspecies (Banff longnose dace) is now extinct. Onenative species (chinook salmon) is certainly extirpated in the CCRE. Several otheranadromous species (Pacific lamprey, white sturgeon, sockeye and steelhead stocks)native to the study area in the Columbia River mainstem or adjacent connected waterspresumably are extirpated or landlocked. Many individual stocks of native fish also havebeen extirpated. Cutthroat trout and bull trout native stocks especially have beeneliminated or severely depleted: both are now rare in most of the Bow River drainage. Inparticular, genetically-pure stocks of native westslope cutthroat trout in their nativehabitats are exceedingly rare and should be treated as endangered. A limited number ofcutthroat stocks have been successfully introduced into previously fish-free drainagesabove barriers.

Of the 21 introduced species, at least 13 have managed to persist somewhere within theCCRE. Introduced brook and brown trout have been especially successful, largelyreplacing native cutthroat and bull trout in many east slopes waters. Most rainbow troutstocks introduced into the upper Bow drainage evidently have failed to persist in thelong term, but have introgressed extensively with native westslope cutthroats, especiallyin the Bow mainstem, Jumpingpound Creek, Elbow, Sheep and Highwood riverdrainages. Introduced lake trout stocks have replaced native bull trout in at least twoBNP lakes even though certain native stocks of these two species coexist elsewhere in theCCRE. An introduced stock of northern Dolly Varden may pose a threat to a native bulltrout stock being restored in the upper Kananaskis drainage. One introduced stock (ciscoin Lake Minnewanka) derived from the Great Lakes may be of special value forconservation because stocks in lakes Ontario and Erie are thought to be endangered.

The present fish fauna of the CCRE is overwhelmingly artificial, depauperate anddominated by exotic stocks; the native fish fauna largely obliterated. The ecologicalintegrity of the natural Central Rockies Ecosystem has been irretrievably lost. A system oftriage is required to salvage whatever remains of the natural state. A new set ofecosystem boundaries based on regional watersheds is proposed within which toorganize aquatic ecosystem conservation and management in the future.

Acknowledgements

Numerous locality records for fishes were provided by Parks Canada,Banff National Park (BNP); the Alberta Fish and Wildlife Service (AFW),Calgary and Rocky Mountain House; and the British Columbia Ministry ofEnvironment, Lands and Parks (BCME), Cranbrook. Marnie Pole providedsome locality records for Yoho National Park. Charlie Pacas, Tom Hurdand John Paczkowski contributed some recent locality records for theClearwater, Red Deer, lower Spray and Cascade rivers in BNP. Jim Stelfox,Mel Kraft, Larry Rhude and Steve Herman of AFW allowed me to usereports, distribution maps and file data for waters in their jurisdiction,while Bill Westover of BCME did the same for the Columbia andKootenay drainages in British Columbia. All of the above provided muchuseful discussion, corrections and additions based on their localknowledge. Ray Blanchard and Jim Rennels directed me to certaincutthroat trout records. Charlie Pacas and Larry Rhude provided helpfulcomments on an earlier draft. Part of this study was funded by CanadianParks Service contract KBP-2059, managed for BNP by Tom Hurd (1992-93) and Charlie Pacas (1993-94). This report is submitted in partialfulfillment of that contract.

Table of Contents

Introduction . . . . . . . . . . . . . . . . . . . 1

Methods . . . . . . . . . . . . . . . . . . . 3

The Fishes . . . . . . . . . . . . . . . . . . . 6Pacific lamprey . . . . . . . . . . . . . . . . 6Lake sturgeon . . . . . . . . . . . . . . . . 10White sturgeon . . . . . . . . . . . . . . . . 11Chiselmouth . . . . . . . . . . . . . . . . . 11Lake chub . . . . . . . . . . . . . . . . . . 11Brassy minnow . . . . . . . . . . . . . . . . 12Pearl dace . . . . . . . . . . . . . . . . . . 12Peamouth . . . . . . . . . . . . . . . . . . 12Northern redbelly dace . . . . . . . . . . . . . 12Finescale dace . . . . . . . . . . . . . . . . 12Fathead minnow . . . . . . . . . . . . . . . 13Northern squawfish . . . . . . . . . . . . . . 13Blacknose dace . . . . . . . . . . . . . . . . 13Longnose dace . . . . . . . . . . . . . . . . 13Leopard dace . . . . . . . . . . . . . . . . . 14Redside shiner . . . . . . . . . . . . . . . . 14Longnose sucker . . . . . . . . . . . . . . . 14White sucker . . . . . . . . . . . . . . . . . 15Largescale sucker . . . . . . . . . . . . . . . 15Mountain sucker . . . . . . . . . . . . . . . 15Northern pike . . . . . . . . . . . . . . . . 15Golden trout . . . . . . . . . . . . . . . . . 16Cutthroat trout . . . . . . . . . . . . . . . . 16Rainbow trout . . . . . . . . . . . . . . . . 22Coho salmon . . . . . . . . . . . . . . . . . 22Sockeye salmon & kokanee . . . . . . . . . . . . 22Chinook salmon . . . . . . . . . . . . . . . . 24Atlantic salmon . . . . . . . . . . . . . . . . 24Brown trout . . . . . . . . . . . . . . . . . 25Arctic charr . . . . . . . . . . . . . . . . . 25Bull trout . . . . . . . . . . . . . . . . . . 25

Brook trout . . . . . . . . . . . . . . . . . . 28Dolly Varden . . . . . . . . . . . . . . . . . 31Lake trout . . . . . . . . . . . . . . . . . . 31Splake . . . . . . . . . . . . . . . . . . . 32Cisco . . . . . . . . . . . . . . . . . . . 33Lake whitefish . . . . . . . . . . . . . . . . 33Pygmy whitefish . . . . . . . . . . . . . . . . 33Mountain whitefish . . . . . . . . . . . . . . . 34Arctic grayling . . . . . . . . . . . . . . . . 34Trout-perch . . . . . . . . . . . . . . . . . 34Burbot . . . . . . . . . . . . . . . . . . . 34Western mosquitofish . . . . . . . . . . . . . . 34Sailfin molly . . . . . . . . . . . . . . . . . 36Guppy . . . . . . . . . . . . . . . . . . . 36Green swordtail . . . . . . . . . . . . . . . . 36Brook stickleback . . . . . . . . . . . . . . . 36Mottled sculpin . . . . . . . . . . . . . . . . 36Slimy sculpin . . . . . . . . . . . . . . . . . 37Torrent sculpin . . . . . . . . . . . . . . . . 37Spoonhead sculpin . . . . . . . . . . . . . . . 37Sunfish (unidentified) . . . . . . . . . . . . . . 37Smallmouth bass . . . . . . . . . . . . . . . . 37Largemouth bass . . . . . . . . . . . . . . . . 38Convict cichlid . . . . . . . . . . . . . . . . 38African jewelfish . . . . . . . . . . . . . . . . 38Freshwater angelfish . . . . . . . . . . . . . . 38

DiscussionFaunal Loss: Causes and Consequences . . . . . . . . 39

Westslope Cutthroat Trout . . . . . . . . . . 39Bull Trout . . . . . . . . . . . . . . . . 43Other Species . . . . . . . . . . . . . . 44

The New Fauna . . . . . . . . . . . . . . . . 45Realistic Response:

Ecosystems for Fish Conservation & Management . . 46

References Cited . . . . . . . . . . . . . . . . . . 51

Figures & Tables

Figure 1: The study area . . . . . . . . . . . . . . . 2Figure 2: Westslope cutthroat trout native distribution . . . . . 17Figure 3: Westslope cutthroat trout present distribution . . . . 20Figure 4: Rainbow trout distribution . . . . . . . . . . . 23Figure 5: Brown trout distribution . . . . . . . . . . . . 26Figure 6: Bull trout native distribution . . . . . . . . . . 27Figure 7: Bull trout present distribution . . . . . . . . . . 29Figure 8: Brook trout distribution . . . . . . . . . . . . 30Figure 9: Mountain whitefish native distribution . . . . . . . 35Figure 10: Proposed major regional watershed ecosystems . . . 50

Table 1: Scientific & common names of fishes . . . . . . . . 7Table 2: Fishes of the CCRE . . . . . . . . . . . . . . 9Table 3: Status of native fishes . . . . . . . . . . . . . 40Table 4: Status of introduced fishes . . . . . . . . . . . 45

Introduction

Introduction

Banff National Park (BNP) engaged Freshwater Research in September1992

1. to prepare a fish management plan for BNP,

2. to conduct a preliminary fall survey of the fishes of the Bow River fromabove Bow Falls to the headwaters, and

3. to prepare a chapter for the Central Canadian Rockies Ecosystem(CCRE) Atlas on the distribution and status of fishes within the CCRE.

This report is an overview of the distribution and status of the fish stockswithin the CCRE. Earlier drafts (1993, 1994) formed the background andsupporting documentation for the chapter in the CCRE Atlas noted underitem (3), above. This final version incorporates the most complete speciesaccounts, distribution maps and interpretation of the data, as well as thedocumentation for the native and present fish distributions.

Yoho, Kootenay and Banff national parks comprise the core of the CCRE,which at the time this study was initiated had the following boundaries.West of the Continental Divide the study area consists of the ColumbiaRiver right bank headwaters from the Bush River upstream, including themainstem Columbia River; the Kootenay River headwaters above CanalFlats; and the Elk River and Fording River headwaters north of Elkford.East of the Continental Divide the study area includes the NorthSaskatchewan, Red Deer and Bow River drainages upstream of the ForestReserve boundary. A very small part of the extreme headwaters of theWillow Creek drainage, Oldman system, are included along the southboundary of the study area. The general locality and precise boundariesare illustrated in Figure 1.

1

Figure 1. The Central CanadianRockies Ecosystem (CCRE), scaleapproximately 1:1,418,440. Basemap Alberta Forestry, Lands andWildlife (1992).

Methods

Methods

This report was prepared entirely from existing data. The distribution andhistorical abundance conclusions represent my interpretations ofinformation in publications, technical reports and selected file data held bythe government agencies responsible for fish management in the threejurisdictions included in the CCRE, interviews with reliable observersknowledgeable about certain waters, and incidental observations in a largenumber of historical records. The specific sources used to establishdistributions of fishes within each major drainage basin are as follows.

General study area: Department of the Interior (1879 to 1918), Sisley(1911), Prince et al. (1912), Whitehouse (1919), Cuerrier and Ward (1952,1953, 1954), Carl et al. (1959), Henderson and Peter (1969), Paetz andNelson (1970), Scott and Crossman (1973), Ward (1974), Mudry andAnderson (1977), Cavender (1978), Lee et al. (1980), Donald (1987), Nelsonand Paetz (1976, 1992), Haas and McPhail (1991)

Columbia River basin: De Smet (1847), Hector (1863), Lees andClutterbuck (1888), Green (1890), Eigenmann and Eigenmann (1892),Eigenmann (1895), Stutfield and Collie (1903), Henshaw (1906), Dymond(1932), McHugh (1940, 1941), Nelson (1968), McCart (1970), Mudry andAnderson (1975), Erasmus (1984), Triton Environmental Consultants Ltd.(1992), Fielden et al. (1992), RL&L Environmental Services Ltd. (1993), M.Pole (CPS Field, BC, personal communication 1991), W. Westover (BCMECranbrook, BC, interview 1993)

Kootenay River basin: Lees and Clutterbuck (1888), Hornaday (1907),Dymond (1932), Nelson (1968), McAllister et al. (1981), Alger and Donald(1984), Donald and Alger (1984), SHIP Environmental Consultants Ltd.(1991), B. Sheehan (Warden Service, Kootenay National Park, BC,interview 1993), W. Westover (BCME Cranbrook, BC, interview 1993)

North Saskatchewan River basin: Stutfield and Collie (1903), Sch�ffer(1911), Miller and Paetz (1953), Tebby (1974); Allan (1980); Donald andAlger (1993); AFW summary file data mapped to 1993; S. Herman, M.Kraft and L. Rhude (AFW Rocky Mountain House, interviews 1993)

3

Methods

Red Deer River basin: Miller and Paetz (1953), Anderson and Donald(1978), AFW summary file data mapped to 1993; S. Herman, M. Kraft andL. Rhude (AFW Rocky Mountain House, interviews 1993)

Bow River basin: De Smet (1847); Hector (1863); Palliser (1863); Whitcher(1887); Eigenmann and Eigenmann (1892); Eigenmann (1895); Wilcox(1896, 1900); Stutfield (1899); Stutfield and Collie (1903); Macoun (1905);Outram (1905); Vick (1913); Rawson (1939, 1942, 1945, 1947); McHugh(1940, 1941); Edwards (1950); Miller and Macdonald (1950); Cuerrier(1954); Cunningham (1961, 1962, 1964); Nelson (1965); McAllister (1969);Thompson (1975, 1977, 1978); Davies and Thompson (1976); Howes (1976);Mayhood and Anderson (1976); Mayhood et al. (1976); Mudry and Green(1976, 1977); Thompson and Davies (1976); Anderson and Donald (1978);Mayhood (1978, 1983, 1985); Wiebe (1978); Tripp et al. (1979); Donald et al.(1980); Techman Ltd. (1980); Stelfox and Konynenbelt (1980); McAllister etal. (1981); Monenco Consultants Limited (1982); Stelfox and Ladd (1982);Lamoureux et al. (1983); McIllrie and White-Fraser (1983); Nelson (1983);Stelfox (1983, 1985); Nibourg (1985); Matkowski and Fernet (1987); Carland Stelfox (1989); Colpitts (1991); Hills and Stelfox (1991); Edworthy(1992); Sentar (1992); Donald and Alger (1993); Mayhood and Paczkowski(1993); J. Stelfox (AFW Calgary, AB, interviews to 1994); T. Hurd, C. Pacasand J. Paczkowski (CPS Banff, AB, interviews to 1994).

The historical documents were useful for establishing the native presenceof trout and salmon in many waters. Although there were some importantexceptions, seldom were historical descriptions clear enough to identifyspecies; nor were the authorsÕ own identifications reliable enough toaccept at face value in most cases. Detailed analyses of technical andhistorical documents upon which much of the present interpretation ofnative trout distributions in the Rocky Mountains is based are currentlybeing prepared and will be presented elsewhere.

In general I obtained the most complete and reliable information for theBow drainage, and for Yoho and Kootenay national parks in the Columbiaand Kootenay drainages, respectively. Additional primary material inextensive agency files remains to be included for all basins, especially theNorth Saskatchewan and Red Deer drainages under the jurisdiction of theProvince of Alberta, and the Columbia and Kootenay drainages under thejurisdiction of the Province of British Columbia. Existing data for the

4

Methods

Kootenay basin within BC provincial jurisdiction appear to extend somefish distributions far downstream based on upstream occurrence records;actual distributions in that drainage may be much less continuous thanshown on the distribution maps. Ongoing surveys of the Columbiawatershed under the Mica Dam Compensation Program are adding newinformation for the latter drainage, the northern part of which has beenchanged dramatically by the closure of the Mica Dam.

5

The Fishes

The Fishes

Scientific and common names of the fishes mentioned in this report arelisted in Table 1. The distribution of fishes in the CCRE is summarized bymajor drainage basin in Table 2. The detailed distributions of severalspecies for which substantial data were available are illustrated in separatefigures.

Pacific lamprey Ñ Carl et al. (1959) gave the distribution of this species asincluding Òthe headwaters of the Columbia.Ó There is a specimen recordfor the Columbia River apparently near the Bush River in the CCRE (Scottand Crossman 1973:43). There have been numerous recent changes in thehabitat here due to damming, and the present status of this fish in theCCRE is in doubt: lampreys have not been reported by recent investigatorsin this area (Fielden et al. 1992, Triton Environmental Consultants Ltd.1992).

An unusual feature of this fish is that it is able to use its sucker to scalewaterfalls, dams and other impediments that are barriers to other fishes(Carl et al. 1959:27, Scott and Crossman 1973:43). There surely must besome doubt, however, whether it is capable of ascending the 13 dams onthe Columbia mainstem, some among the largest in the world, over thenearly 1500-km distance from the Pacific to Kinbasket reservoir. Whetherthe population in the upper Columbia is a strictly freshwater form was notreported. The conservation status of the species has not been investigatedin Canada (Campbell 1993:396-397). The conservation status in NorthAmerica is unlisted, except for one as yet undescribed subspecies ofspecial concern in California and Oregon (Williams et al. 1989:3).

Beamish (1982) described a new species, the Vancouver lamprey, which issimilar to the Pacific lamprey and easily mistaken for it. The new species isa freshwater form, whereas the Pacific lamprey ostensibly is obligatelyanadromous. The Vancouver lamprey is presently listed as vulnerable byCOSEWIC (Campbell 1993:396, as lake lamprey). It is therefore ofconsiderable interest to know whether a lamprey population still exists inthe CCRE, and if so, if it is truly the Pacific lamprey, the new species, or asimilar but as yet undescribed form.

6

The Fishes

Table 1. Scientific and common names of fishes mentioned in this report (Robins 1991a, 1991b). Subspeciesnames from Renaud and McAllister (1988) and Behnke (1992). The spelling of charr follows Morton (1980).

Petromyzontidae - lampreysVancouver lamprey Lampetra macrostoma Beamish, 1982Pacific lamprey Lampetra tridentata (Gairdner, 1836)

Acipenseridae - sturgeonslake sturgeon Acipenser fulvescens Rafinesque, 1817white sturgeon Acipenser transmontanus Richardson, 1836

Cyprinidae - minnowschiselmouth Acrocheilus alutaceus Agassiz & Pickering, 1855lake chub Couesius plumbeus (Agassiz, 1850)brassy minnow Hybognathus hankinsoni Hubbs, 1929pearl dace Margariscus margarita Cope, 1868peamouth Mylocheilus caurinus (Richardson, 1836)northern redbelly dace Phoxinus eos (Cope, 1862)finescale dace Phoxinus neogaeus Cope, 1868fathead minnow Pimephales promelas Rafinesque, 1820northern squawfish Ptychocheilus oregonensis (Richardson, 1836)blacknose dace Rhinichthys atratulus (Hermann, 1804)longnose dace Rhinichthys cataractae (Valenciennes, 1842)

Banff longnose dace R. c. smithi Nichols, 1916leopard dace Rhinichthys falcatus (Eigenmann & Eigenmann, 1893)redside shiner Richardsonius balteatus (Richardson, 1836)

Catostomidae - suckerslongnose sucker Catostomus catostomus (Forster, 1773)white sucker Catostomus commersoni (Lacepède, 1803)largescale sucker Catostomus macrocheilus Girard, 1856mountain sucker Catostomus platyrhynchus (Cope, 1874)

Esocidae - pikesnorthern pike Esox lucius Linnaeus, 1758

Salmonidae - salmons, trouts, charrs, whitefishes & graylingswhitefishescisco Coregonus artedi Lesueur, 1818lake whitefish Coregonus clupeaformis (Mitchill, 1818)pygmy whitefish Prosopium coulteri (Eigenmann & Eigenmann, 1892)mountain whitefish Prosopium williamsoni (Girard, 1856)western troutsgolden trout Oncorhynchus aguabonita (Jordan, 1893)cutthroat trout Oncorhynchus clarki (Richardson, 1836)

Yellowstone O. c. bouvieri (Bendire, 1882)coastal O. c. clarki (Richardson, 1836)westslope O. c. lewisi (Girard, 1856)

rainbow trout Oncorhynchus mykiss (Walbaum, 1792)

continued …

7

The Fishes

Table 1. concluded

Pacific salmonscoho salmon Oncorhynchus kisutch (Walbaum, 1792)sockeye salmon Oncorhynchus nerka (Walbaum, 1792)chinook salmon Oncorhynchus tshawytscha (Walbaum, 1792)eastern trout and salmonAtlantic salmon Salmo salar Linnaeus, 1758brown trout Salmo trutta Linnaeus, 1758charrsArctic charr Salvelinus alpinus (Linnaeus, 1758)bull trout Salvelinus confluentus (Suckley, 1858)brook trout Salvelinus fontinalis (Mitchill, 1814)Dolly Varden Salvelinus malma (Walbaum, 1792)lake trout Salvelinus namaycush (Walbaum, 1792)splake Salvelinus fontinalis × S. namaycushgraylingsArctic grayling Thymallus arcticus (Pallas, 1776)

Percopsidae - trout-perchestrout-perch Percopsis omiscomaycus (Walbaum, 1792)

Gadidae - codsburbot Lota lota (Linnaeus, 1758)

Poeciliidae - live-bearerswestern mosquitofish Gambusia affinis (Baird & Girard, 1853)sailfin molly Poecilia latipinna (Lesueur, 1821)guppy Poecilia reticulata Peters, 1859green swordtail Xiphophorus helleri Heckel, 1848

Gasterosteidae - sticklebacksbrook stickleback Culaea inconstans (Kirtland, 1841)

Cottidae - sculpinsmottled sculpin Cottus bairdi Girard, 1850slimy sculpin Cottus cognatus Richardson, 1836torrent sculpin Cottus rhotheus (Smith, 1882)spoonhead sculpin Cottus ricei (Nelson, 1876)

Centrarchidae - sunfishessunfish (unidentified) Lepomis sp.smallmouth bass Micropterus dolomieu Lacepède 1802largemouth bass Micropterus salmoides (Lacepède 1802)

Cichlidae - cichlidsconvict cichlid Cichlasoma nigrofasciatum (Günther, 1867)African jewelfish Hemichromis bimaculatus Gill, 1862freshwater angelfish Pterophyllum scalare (Lichtenstein, 1823)

8

The Fishes

Table 2. The fishes of the Central Canadian Rockies Ecosystem. ● - native, ❍ - native in adjacent connectedwaters, I - introduced, U - unsuccessful introduction, ? - questionable record. Some species and stocks havebeen extirpated, and many others are at risk. The Kicking Horse records refer only to occurrences aboveWapta Falls.

NorthRed Sask- Kicking

Species Bow Deer atchewan Columbia Horse Kootenay

Petromyzontidae - lampreysPacific lamprey ●

Acipenseridae - sturgeonslake sturgeon ❍? ❍white sturgeon ❍

Cyprinidae - minnowslake chub ● ● ● ?pearl dace ● ● ●northern redbelly dace ●finescale dace ●fathead minnow ❍ ● ●longnose dace ● ● ● ● ●chiselmouth ●peamouth ●northern squawfish ● ❍leopard dace ❍?redside shiner ● ❍

Catostomidae - suckerslongnose sucker ● ● ● ● ●? ●white sucker ● ● ●mountain sucker ● ● ●largescale sucker ● ●? ?

Esocidae - pikesnorthern pike ❍ ● ●

Salmonidae - salmons, trouts, charrs, whitefishes & graylingschinook salmon ●sockeye salmon ❍?kokanee I Icoho salmon I?golden trout U I U Ucutthroat trout ● I I ● ●? ●rainbow trout I I I ● ●? Ibrown trout I I IAtlantic salmon U Ubull trout ● ● ● ● ● ●brook trout I I I I I IDolly Varden Ilake trout ● ● IArctic charr Usplake I U U Ucisco Ilake whitefish I?

continued …

9

The Fishes

Table 2. concluded

NorthSask- Kicking

Species Bow Red Deer atchewan Columbia Horse Kootenay

pygmy whitefish ● ●mountain whitefish ● ● ● ● ● ●

Percopsidae - trout-perchestrout-perch ● ●

Gadidae - codsburbot ● ● ● ●

Poeciliidae - live-bearerswestern mosquitofish Isailfin molly Iguppy Ugreen swordtail U

Gasterosteidae - sticklebacksbrook stickleback ● ●

Cottidae -sculpinsspoonhead sculpin ❍ ● ●slimy sculpin ● ● ●torrent sculpin ❍ ● ❍mottled sculpin ●?

Centrarchidae - sunfishessunfish unidentified Ilargemouth bass U? Ismallmouth bass U?

Cichlidae - cichlidsAfrican jewelfish Iconvict cichlid Ufreshwater angelfish U

Lake sturgeon Ñ There is a 1970 specimen record as near to the CCRE asthe Clearwater River (NorthÊSaskatchewan drainage) betweenChedderville and Ricinus, according to Nelson and Paetz (1976), citing apersonal communication with W. E. Roberts. Writing in 1890, J. H. McIllrie(McIllrie and White-Fraser 1983) reported that Òa few sturgeon come up[rivers in the Calgary district] until the water gets too rapid for themÓ.Whether they reached the CCRE in the Bow, Elbow or Highwood is notclear: there are no records, but there are no impassable natural obstacleseither. Dams now prevent them from from ascending the Bow drainageabove the extreme lower reach of the Bow River. The Dickson dam on theRed Deer presently prevents movements of lake sturgeon into the CCREin that drainage, unless a population has become established in thereservoir. The species is considered threatened throughout North America(Williams et al. 1989:3). Although COSEWIC currently lists it as Òreport

10

The Fishes

accepted, no status designation requiredÓ in Canada, this designation ispresently under review (Campbell 1993:396, 398). Rochard et al. (1990:131)concluded that all species of sturgeon can be considered endangered,based on considerations of life histories and threats to existence for 24species worldwide.

White sturgeon Ñ Carl et al. (1959) showed the distribution of this fish asincluding the entire upper Columbia River drainage, but the closest actualspecimen record to the CCRE is for the Columbia River above Revelstoke(Scott and Crossman 1973:97). Its present status in the CCRE is unknown.Its conservation status in North America is unlisted (Williams et al. 1989),but the genetically-distinct Kootenai (Kootenay) River population is beingconsidered for listing as endangered under the Endangered Species Act inthe United States (Duke 1993). The species is listed as vulnerable byCOSEWIC (Campbell 1993:396). Rochard et al. (1990:131) concluded thatall species of sturgeon can be considered endangered, based onconsiderations of life histories and threats to existence for 24 speciesworldwide.

Chiselmouth Ñ There is one record of this minnow within the CCRE:from Windermere Lake, on the Columbia River mainstem (Carl et al.1959:119). The species is known in Canada only from a few widely-separated localities, all of them in the Fraser and Columbia systems ofBritish Columbia. The chiselmouth presently is under review byCOSEWIC for possible listing as vulnerable in British Columbia (Campbell1993:398).

Lake chub Ñ Lake chub are widespread in the CCRE in the NorthSaskatchewan, Red Deer and Bow drainages. They are said to exist also inthe east Kootenay region of the Columbia system (Carl et al. 1959:122), butthere appears to be only a single published locality record for that area Ñnear Cranbrook, south of the CCRE (Scott and Crossman 1973:403, Lee etal. 1980:150). The species has not been reported in recent collections in theEast Kootenay portion of the study area (Alger and Donald 1984, Donaldand Alger 1984, Fielden et al. 1992, Triton Environmental Consultants Ltd.1992, RL & L Environmental Services Ltd. 1993). Lake chub have recentlybeen discovered in Dog Lake, Kootenay drainage, Kootenay NationalPark, where they are believed to have been illegally introduced (B.Sheehan, personal communication 1993; cf. Donald and Alger 1984).Introduced stocks in the profoundly disrupted habitats of reservoirs are

11

The Fishes

known to hybridize with introduced longnose dace (Nelson 1966).

Lake chub east and west of the Continental Divide likely represent twodistinct geographic stocks, possibly separate subspecies (McPhail andLindsey 1970:244-5).

Brassy minnow Ñ A specimen record for this species in the Bow Riverdrainage in or near the CCRE (Lee et al. 1980:175) undoubtedly iserroneous (cf. Nelson and Paetz 1992:133). There is no indisputable recordof the brassy minnow within the CCRE.

Pearl dace Ñ Pearl dace have been recorded from the CCRE in theheadwaters of Shunda Creek, North Saskatchewan drainage (Nelson andPaetz 1992), from James River, Red Deer drainage (Henderson and Peter1969:330), and from Little Jumpingpound Creek (Nibourg 1985), all on theextreme eastern edge of the study area. Pearl dace also have been recordedfrom the Bow drainage close to the CCRE in the Sheep River watershed(Nelson and Paetz 1992). They are to be expected in suitable habitat (boggywaters lacking piscivorous fish) east of the mountain front throughout thestudy area.

Peamouth Ñ This species is said to be widespread in the upper Columbiasystem (Carl et al. 1959:112, Scott and Crossman 1973:425-6). There arecollection records for peamouth in the CCRE at Golden (Eigenmann 1895)and in Columbia Lake on the Columbia mainstem (Scott and Crossman1973:425, Lee et al. 1980:208, RL & L Environmental Services Ltd. 1993).

Northern redbelly dace Ñ There is a single record of this species in theCCRE, from the North Saskatchewan drainage near Nordegg, at theextreme northeastern edge of the study area (Nelson and Paetz 1992:152).This and the following species commonly hybridize. The hybrid mayconstitute a separate single-sex (all female) species that requires males ofone of the parental species for successful clonal reproduction (Dawley etal. 1987, Goddard and Dawley 1990, Nelson and Paetz 1992:153-154). Thehybrid should be carefully sought in the CCRE wherever either parentalspecies is found.

Finescale dace Ñ There is a single record of this species in the CCRE,from the North Saskatchewan drainage near Nordegg, at the extreme

12

The Fishes

northeastern edge of the study area (Nelson and Paetz 1992:156). See thediscussion of hybridization of this species with northern redbelly dace inthe above entry.

Fathead minnow Ñ Fathead minnows have been captured from twolocations within the CCRE: near Nordegg, in the North Saskatchewandrainage, and from the James River drainage in the Red Deer system(Nelson and Paetz 1992:161). The species also occurs in Bighill Creek, aBow River tributary a few miles below the CCRE (Nelson 1965:744).

Northern squawfish Ñ Squawfish are widespread in the upper Columbiasystem, where the species has been recorded within the CCRE in theColumbia River mainstem, and near the CCRE in the Kootenay drainage(Carl et al. 1959, Lee et al. 1980:349). There are no records of the species inthe Kicking Horse watershed above Wapta Falls, however (M. J. Pole,personal communication).

Blacknose dace Ñ Rawson (1939:12, 1942:12) reported this species (asRhinichthys atronasus) in collections from Lake Minnewanka, BNP (Bowdrainage). Nelson and Paetz (1992:379) suggest that these fish were mostlikely hybrids of lake chub and longnose dace. Blacknose dace, an easternspecies, are otherwise unknown in either British Columbia or Albertawaters (Lee et al. 1980:352).

Longnose dace Ñ This minnow is widespread within the CCRE in theBow and North Saskatchewan drainages, but has been reported only in theJames River drainage in the Red Deer basin within the study area(Henderson and Peter 1969:330, Nelson and Paetz 1992:171). There areseveral locality records above impassable falls in the Ram River drainage,North Saskatchewan system (Henderson and Peter 1969:330; Paetz andNelson 1970:132; Nelson and Paetz 1992:171, 172; L. Rhude, Alberta Fishand Wildlife Service, personal communication). Although these raise astrong suspicion of illicit introduction by bait bucket, they could in fact benative: detailed morphological, genetic and distributional studies areneeded. The species is said to be widespread in the Kootenay and upperColumbia drainages (Carl et al. 1959:128). There are recent specimenrecords from the Columbia mainstem drainage, one within the CCRE inSuccour Creek, tributary to the Bush River (Fielden et al. 1992, TritonConsulting Services Ltd. 1992). This species has not been recorded fromthe Kicking Horse drainage above Wapta Falls (M. J. Pole, personal

13

The Fishes

communication 1991). Introduced longnose dace hybridize withintroduced lake chub in the disrupted habitats of mountain reservoirs(Nelson 1966, Nelson and Paetz 1992:379).

The Banff longnose dace (R. c. smithi), an endemic subspecies describedfrom Cave and Basin Hotsprings in BNP by Nichols (1916), is now extinct(Renaud and McAllister 1988, Miller et al. 1989, Campbell 1993:397).

Leopard dace Ñ The single record of this species anywhere near theCCRE, in the Kootenay drainage near Cranbrook (Carl et al. 1959:120, Scottand Crossman 1973, Lee et al. 1980), was considered questionable byPeden (1991:181). Peden (1991) believes Columbia River populations inBritish Columbia are vulnerable, mainly as a result of extensivehydroelectric development.

Redside shiner Ñ Carl et al. (1959:106) indicate that the redside shiner isgenerally distributed throughout the upper Columbia system, includingthe headwaters of the Kootenay River. The species has been recordedwithin the CCRE from the Columbia River mainstem tributaries andColumbia Lake (Eigenmann 1895, Triton Consulting Services Ltd. 1992, RL& L Environmental Services Ltd. 1993). A published specimen recordexists for the Kootenay drainage just south of the CCRE (Lee et al.1980:358).

Longnose sucker Ñ The longnose sucker is widespread in the CCRE inthe North Saskatchewan and Bow drainages, though curiously it has notbeen recorded further upstream than the CCRE eastern boundary in theJames River drainage of the Red Deer basin (Nelson and Paetz 1992:191). Arecently-discovered population beside a major road above impassable fallson the Ram River, North Saskatchewan drainage (L. Rhude, Alberta Fishand Wildlife Service, personal communication) requires morphologicaland genetic study to determine if it is native or an illicit introduction. Thespecies is said to occur throughout the Columbia River system in BC (Carlet al. 1959:91). Specimen records exist for the Columbia mainstem nearKinbasket Lake below the CCRE (Scott and Crossman 1973:533, Lee et al.1980:372), and within the CCRE at Golden (Eigenmann 1895) and SuccourCreek (Fielden et al. 1992). RL & L Environmental Services Ltd. (1993)reported capturing a single specimen from Columbia Lake. The species isknown from the Kootenay drainage within the study area in Daer Pond

14

The Fishes

and Dog Lake, KNP (Donald and Alger 1984, Donald 1987), where it ispresumed to be native. It has been suggested that longnose suckers in theKicking Horse drainage above Wapta Falls are not native there (M. J. Pole,personal communication). Introduced longnose suckers hybridize withintroduced white suckers in certain mountain hydroelectric reservoirs inthe CCRE (Nelson 1973).

White sucker Ñ White suckers have been reported from the NorthSaskatchewan, Red Deer and Bow drainages within the CCRE (Hendersonand Peter 1969:331, Nelson and Paetz 1992:194). West of the ContinentalDivide within the study area this species is absent, being replaced by thelargescale sucker (Carl et al. 1959:87, Lee et al. 1980). White suckers in theupper Kicking Horse drainage (M. Pole, personal communication 1991) areundoubtedly introduced. White suckers evidently have expanded theirrange in the Bow drainage to above Bow Falls, probably as a result ofhuman intervention (Sentar 1992, Mayhood and Paczkowski 1993).Introduced white suckers hybridize with introduced longnose suckers inthe highly disturbed habitats provided by some mountain hydroelectricreservoirs (Nelson 1973).

Largescale sucker Ñ Carl et al. (1959:87) indicate by map shading thatlargescale suckers are found throughout the Columbia and Kootenaydrainages. Recent specimen records have been reported in the CCRE forColumbia Lake (RL & L Environmental Services Ltd. 1993). TritonEnvironmental Consultants Ltd. (1992) reported largescale suckers in theircatches from upper Columbia River tributaries just outside the CCRE.Largescale suckers in Ottertail River, in Yoho National Park above WaptaFalls, are believed to have been introduced (M. Pole, personalcommunication 1991).

Mountain sucker Ñ The mountain sucker is absent from the CCRE westof the Continental Divide (Carl et al. 1959, Lee et al. 1980), but has beenfound in the study area east of the divide in the North Saskatchewan, RedDeer and Bow drainages (Henderson and Peter 1969:331, Nelson andPaetz 1992:200).

Northern pike Ñ Pike have been found in the CCRE the James Riverdrainage of the Red Deer basin (Henderson and Peter 1969:329). Thespecies is also known from Abraham Reservoir and Swan Lake, NorthSaskatchewan drainage (Nelson and Paetz 1992:219, Alberta Fish and

15

The Fishes

Wildlife Service file data). Pike occur elsewhere in the NorthSaskatchewan drainage downstream from the study area (Henderson andPeter 1969:329, Nelson and Paetz 1992:218). Monenco Consultants Limited(1982) reported a single specimen from lower Threepoint Creek, justdownstream from the CCRE in the Bow system. Conceivably the speciesoccasionally enters the study area in this drainage. Pike are not native tothe Columbia system, occurring there only as introduced populations in afew locations far downstream in the USA (Scott and Crossman 1973, Lee etal. 1980).

Golden trout Ñ Attempts to establish non-native golden trout in CCREwaters have been made in Banff, Kootenay and Yoho national parks (Ward1974; B. Sheehan, personal communication 1993), and in the NorthSaskatchewan (Coral, Gap, Michelle lakes), Red Deer (Lost Guide Lake)and Bow (Three Isle and Galatea lakes, Kananaskis drainage) systemsunder Alberta provincial jurisdiction (Nelson and Paetz 1992:272, AlbertaFish and Wildlife Service file data). Of these, only Coral Lake still holdsthe species (Ward 1974; Nelson and Paetz 1992; L. Rhude, Alberta Fish andWildlife Service, personal communication). One stock in California isthreatened (Williams et al. 1989).

Cutthroat trout Ñ The probable native distribution of westslopecutthroat trout in the CCRE is presented in Figure 2. In interpreting theexisting record, I have usually used barriers to migration where these areknown to me to set the upstream limits of native cutthroat distributions. Ingeneral I have assumed that waters with apparently suitable cutthroathabitat that are or were accessible to known native stocks were once usedby those stocks.

Figure 2 is necessarily somewhat speculative, representing myinterpretation of numerous disparate technical reports and many dozensof sometimes confusing historical records. Early observers were clearlybewildered by the many apparent varieties of trout, knew them bynumerous undescriptive and apparently interchangeable common names,and rarely described the types in sufficient detail to be identifiable bypresent-day biologists. In fact the cutthroat trout often was notdistinguished from rainbow trout Ñ even by biologists Ñ in the last halfof the nineteenth century (Eigenmann 1895, Macoun 1905, Scott andCrossman 1973:182-3, cf. 191). The species was not given the common

16

Figure 2. Native distribution ofwestslope cutthroat trout in the CCRE.

The Fishes

name ÒcutthroatÓ until 1884 (Behnke 1992:53), and certainly did notbecome widely known by it until much later (cf. Macoun 1905). Thushistorical records of Òrainbow troutÓ frequently referred to cutthroat trout,as indeed angler reports often do even today.

Westslope cutthroat trout are native to the CCRE in the Bow and Kootenaysystems (Behnke 1992:78), and appear to have been native to certainColumbia River headwater tributaries in the vicinity of Columbia andWindermere lakes (e.g., Dutch and Toby creeks, Prince et al. 1912:38). Thespecies probably is native to several other Columbia River tributariesupstream of Brisco, where it is the only black-spotted trout occupyingseveral left bank tributaries according to one recent anglerÕs report(Ambrosi 1987). Few of these have been shown on the map because I havebeen unable to find documentary evidence of cutthroats in most right-bank streams in that area. Elsewhere in the upper Columbia system thepredominant native black-spotted trout is the rainbow, and nativewestslope cutthroats exist (or once existed) only as disjunct relictpopulations in a half-dozen locations, usually above barrier falls (Dymond1932, Carl et al. 1959, Behnke 1992).

One of these disjunct stocks may have been native to the Kicking Horsedrainage of Yoho National Park above Wapta Falls. ÒTroutÓ were nativeabove Wapta Falls (Hector 1863, cf. Erasmus 1984), Òthree kindsÓapparently providing good fishing in or near Emerald Lake in 1892(McArthur 1893). One, a Òsmall grey troutÓ (bull trout?), was abundant inthe lake in 1887 (McArthur 1888). ÒRainbow troutÓ were common in theEmerald Lake area and possibly Porcupine Creek at an early date,attaining sizes of over 6 pounds (Henshaw 1906). Henshaw included inher article a photograph of a square-tailed ÒrainbowÓ of over twelvepounds from an undisclosed location. After noting that Òrainbow trout orcut-throat trout (Salmo mykiss)Ó was the black-spotted trout of the Rockymountain waters of the Bow and Oldman drainages, Macoun (1905)reported that Òpassing the summitÓ of the Rockies and descending to theColumbia, Òanother black-spotted species is met with, named Steel-heador GairdnerÕs trout (Salmo gairdneri).Ó It is not clear whether thisstatement is intended to mean that S. gairdneri is found in addition to orin place of S. mykiss, but the former alternative would account forMcArthurÕs third kind of trout. I have mapped cutthroat trout as nativeabove Wapta Falls in the Kicking Horse drainage so that due attention will

18

The Fishes

be given to conserving what may be a rare, possibly unique native stock.

The westslope cutthroat has been widely transplanted within its nativeBow and Kootenay systems, as well as into the Red Deer, NorthSaskatchewan and Kicking Horse drainages in the CCRE (Rawson 1947,Ward 1974, McAllister et al. 1981, Nelson and Paetz 1992:258). There issome limited evidence of transplantation of cutthroats (subspeciesunknown) into the Columbia mainstem drainage of the CCRE as well. Arainbow/cutthroat hybrid swarm is reported to exist in Whitepine Creek,and there is some minimal evidence of cutthroat genetic introgression intothe Willowbank Creek rainbow population (Triton EnvironmentalConsultants Ltd. 1992). Although these populations may be naturallyintrogressed (given the existence just mentioned of some native cutthroatstocks in the upper Columbia drainage), rainbow and cutthroat troutrarely hybridize where they occur together naturally (Carl et al. 1959:70;Trotter 1991:248; Behnke 1992:80-81, 86).

The earliest recorded introduction of cutthroat trout in the CCRE wasmade in 1915 into Boom, Louise and Moraine lakes (Ward 1974), wherethere appear to have been extant native populations (Wilcox 1900). Earlierunrecorded stocking of exotic fishes, and transplanting of native fishesinto naturally fishless waters, may have begun with the arrival of theCanadian Pacific Railway (CPR) in the mid-1880s in the Bow drainage(Mayhood 1992a) as well as in the upper Kicking Horse drainage, both ofwhich lie along the CPR main line. Nearly all locality records for trout inthese drainages of the CCRE date from after that time, so there will alwaysbe some doubt about the precise limits of the native distributions of fishesin them.

Figure 3 shows the present distribution of cutthroat populations within theCCRE insofar as I have been able to determine it from the recent recordsavailable to me. It shows the distribution of populations of black-spottedtrout bearing the typical red slashmarks under the jaw characteristic ofcutthroat trout; thus it includes any remaining native populations, as wellas introgressed, transplanted and exotic stocks. The distributions in theBow River drainage, the upper Kicking Horse drainage and the Kootenaydrainage within KNP were mapped from actual locality records, and mostof the gaps between them are real. The distributions in the remainingdrainages have been mapped by others to show ranges of distribution;

19

Figure 3. Present distribution of cutthroattrout in the CCRE. All stocks are shown,including transplanted westslope cutthroatstocks, non-native stocks and genetically-introgressed populations still showing someidentifiable cutthroat trout characters.

The Fishes

actual occupancy of these drainages may be much more fragmented thanthat shown.

Even when all nominal cutthroat populations are included, it is obviousfrom a comparison of Figures 2 and 3 that the species has been decimatedthroughout most of its native range in the CCRE. This is especially true inthe Bow drainage, where cutthroat trout of any description are now absentfrom vast reaches of the drainage basin where once they were abundant.The species is virtually extirpated from the mainstem Bow River belowLake Louise, and from nearly all of its tributaries, including the entireKananaskis River and virtually all of its tributaries. Only a fewfragmentary, highly isolated populations remain in the Bow basin.Renowned stocks of westslope cutthroat trout that once supported highquality indigenous and sport fisheries in the Spray and Lower Kananaskislakes eventually disappeared some time after these waters were convertedinto hydroelectric reservoirs and their waters were stocked with rainbowtrout (e.g., Vick 1913, Miller and Macdonald 1950; cf. Nelson 1965; Crosby1990a, 1990b).

The westslope cutthroat trout is officially recognized as a species of specialconcern in Idaho and Montana, but is not so designated in BritishColumbia and Alberta (Johnson 1987). It does not have a conservationstatus designation from the American Fisheries Society (Williams et al.1989) or COSEWIC (Campbell 1993). Nevertheless, uncontaminatedindigenous stocks in their native habitats are extremely scarce at best inAlberta, and need to be immediately recognized as endangered in thatprovince (Mayhood 1989, 1991). This is particularly the case for streamstocks, of which none have been unequivocally identified as genetically-pure natives to date.

The situation for truly native stocks in British Columbia is less clear, butthe wide overlap in the distributions of native cutthroat and introducedrainbow trout, combined with the deliberate production and stocking ofrainbow-cutthroat hybrids in the Kootenay drainage (Dymond 1932, Carlet al. 1959) strongly suggest that introgression is pervasive and pure nativestocks are rare. Any remnant native populations should be flagged as ofexceedingly high value for preservation.

21

The Fishes

Genetically-pure transplanted stocks appear to be widespread in the studyarea. Most of these represent only one or two common genetic lines(Spray-Marvel-Job and Connor stocks) the individual populations ofwhich require no special protection. The remainder, representing isolateddistinctive stocks, should each be treated as endangered due to theiruniqueness, high value for conservation, and their relatively high risk ofextinction due to their isolation.

Rainbow trout Ñ This species is native in the CCRE only to the Columbiamainstem and its direct tributaries (Macoun 1905, Behnke 1992:166). Thereis evidence that rainbows are native to the Kicking Horse drainage aboveWapta Falls (see Cutthroat trout, above). Whether anadromous stocks ofthis species (steelhead) once ascended the Columbia as far as the studyarea is a matter of conjecture, but several other anadromous species did so,and steelhead ascend a similar distance from the sea in the adjacent FraserRiver system (Carl et al. 1959). Barrier falls on the Kootenai (Kootenay)River just below the Libby dam in Montana excluded rainbows from theKootenay drainage in the CCRE (Behnke 1992:171). Rainbow trout havebeen introduced into all major drainages of the study area (Ward 1974,Nelson and Paetz 1992, SHIP Environmental Consultants Ltd. 1991).

The present documented distribution in the study area is shown in Figure4. Populations introduced into lakes in which natural recruitmentprobably is negligible have been omitted. Lake populations of rainbowtrout that reportedly have natural recruitment (Ward 1974:Appendix C)have been included. The present status of most populations requiresconfirmation. Populations in the Bow mainstem within the CCRE, and inthe Kananaskis River, are known to be very sparse (Nelson 1965, Howes1976, Thompson 1977). Dams on the Columbia have extirpated anyanadromous stocks, or have created landlocked populations.

Coho salmon Ñ Lee et al. (1980:94) show a locality record for this speciesin the Kootenay drainage within the CCRE. This portion of the study areais inaccessible to anadromous stocks due to barrier falls in Montana(Behnke 1992:171). If the record is accurate the population must have beenintroduced.

Sockeye salmon & kokanee Ñ Neither anadromous sockeye salmon norkokanee (landlocked sockeye) are believed to be native to the CCRE.

22

Figure 4. The distribution of rainbow troutin the CCRE. All populations exceptperhaps a few in the Columbia drainagehave been introduced. Many populationsare introgressed with cutthroat trout geneswhere the two species occur together.

The Fishes

Kokanee populations are thought to have been derived from anadromousancestors, and anadromous sockeye apparently did not ascend theColumbia much above the Arrow lakes even before the building of thenumerous dams on the mainstem (Nelson 1968). This view, however, iscontradicted by Macoun (1905), who asserted that sockeye ÒÉ are foundin suitable localities É in the Columbia, all around the Big Bend.ÓAlthough Nelson (1968) notes that river piracy (drainage capture) may bethe origin of kokanee in lakes of southeastern BC, the nearest indigenouspopulations to the CCRE mapped by him are in the St. Mary and Moyiedrainages of the Kootenay system, west and southwest of Cranbrook,respectively. Introduced populations now existing in Windermere andColumbia lakes may be derived from introductions into Kinbasketreservoir as part of the Mica dam mitigation work (RL & L EnvironmentalServices Ltd. 1993, W. Westover, BCME, personal communication).Kokanee from Kinbasket reservoir also use for spawning many right banktributaries of the Columbia within the CCRE (Fielden et al. 1992, TritonEnvironmental Consultants Ltd. 1992). In the Kootenay system, recentspawning runs of introduced kokanee reached at least as far upstream asMount Wardle in KNP (Brian Sheehan, KNP Warden Service, personalcommunication). These fish originated in Koocanusa reservoir behindLibby dam (Montana), more than 230 km downstream.

Chinook salmon Ñ Historical records of salmon in the mainstemheadwaters of the Columbia at Windermere and Columbia lakes (e.g., DeSmet 1847, Lees and Clutterbuck 1888), 1900 km from the sea, almostcertainly refer to this species. Eigenmann (1895) identified chinook salmonfrom the Columbia River at Golden. A major spawning area appears tohave been in the gravelly midchannel shallows below Windermere Lake,where the redd-digging activities of these large fish were said to pose asignificant impediment to river navigation (Lees and Clutterbuck1888:102). Numerous dams on the Columbia, especially the Grand Couleedam which was built without any fish passage facilities (Ebel et al.1989:211), have since extirpated these stocks, and the species no longeroccurs in the CCRE.

Atlantic salmon Ñ Atlantic salmon, sometimes identified as thelandlocked form (ouananiche), were introduced into Sawback Creek,Cuthead Lake, the Cascade River, Lake Minnewanka and an isolatedgravel pit in BNP, all in the Bow drainage; and possibly into Sherbrooke

24

The Fishes

Lake in the Kicking Horse drainage. All introductions were apparentlyunsuccessful (Ward 1974, Anderson and Donald 1978, Nelson and Paetz1992, M. J. Pole, personal communication 1991).

Brown trout Ñ This European native has been widely introduced into theBow, Red Deer and North Saskatchewan drainages, where it is largelyconfined to streams of the outer foothills. Within the CCRE it penetratesdeeply into the study area only in the mainstems of the Bow, lowerKananaskis and Clearwater rivers, and the lower reaches of a handful oftheir lower tributaries (Figure 5).

Arctic charr Ñ The marstoni (Quebec red trout) form of this species oncewas introduced, without any subsequent evidence of natural recruitment,into two of the Block lakes and Lake Louise, Bow drainage, and Òin severalareas already containing [bull trout] populationsÓ (Ward 1974). Eggs ofÒred trout,Ó presumably this form, were shipped from Quebec to the Banffhatchery as early as 1914-15, but the disposition of these eggs was notreported (Fisheries Branch 1917:312, cf. 340). The marstoni form isconsidered part of S. a. oquassa, a disjunct, relict subspecies confined tosouthern Quebec, New England and New Brunswick. Certain native NewEngland populations are considered threatened (Williams et al. 1989).COSEWIC currently is reviewing the status of this subspecies in Canada(Campbell 1993:398).

Whether this species is capable of introgressive hybridization with nativebull trout appears to be unknown. If it is, native bull trout populations inthe drainages in which it was introduced may have been contaminatedwith the genes of this species.

Bull trout Ñ The bull trout is native to all major drainages in the CCRE,on both sides of the Continental Divide (Cavender 1978, Haas and McPhail1991). There are many historical records of this fish that provide evidenceof its former great abundance and exceedingly large sizes attainedthroughout the study area.

Figure 6 presents an interpretation of the native range within the CCREbased on historical records, present distribution and stream accessibility.Bull trout have been excluded by impassable falls from extensive portionsof the CCRE. The most notable examples are in the Ram and Siffleur

25

Figure 5. The distribution of browntrout in the CCRE. All populationshave been introduced.

Figure 6. The native distribution ofbull trout in the CCRE.

The Fishes

rivers, North Saskatchewan drainage; and in the Cross, upper Palliser andupper Fording rivers, Kootenay drainage. On the other hand, bull troutappear to be native above several present-day barriers to migration.Examples are Pinto Lake, Cline-North Saskatchewan drainage (Sch�ffer1911:48, Carl et al. 1989); above Wapta Falls in the upper Kicking Horsedrainage, YNP (see Discussion); and in the upper Blaeberry drainageabove Blaeberry Falls, Columbia drainage (Triton EnvironmentalConsultants 1992).

Figure 7 illustrates the present distribution of bull trout in the CCRE asdocumented since 1974, when this fish was last described as generallyabundant in the study area (Ward 1974:8). In many of its native waters thespecies is scarce but still extant (e.g., Bow River drainage above Bow Falls;Mayhood et al. 1976:56-57, Mayhood and Paczkowski 1993). Exceptionsinclude complete losses from Bow and Hector lakes, apparently due todisplacement by lake trout (Donald and Alger 1993), and from SprayLakes after it was converted into a hydroelectric reservoir and stockedwith lake trout (Wilcox 1900, Vick 1913, Miller and Macdonald 1950; cf.Crosby 1990b). Bull trout also are virtually absent from their formerhabitat in the mainstem Bow River below Bow Falls at Banff, and from theKananaskis River mainstem (Nelson 1965, Howes 1976, Thompson 1977).Other apparent differences between the present and native ranges arecertainly due to lack of recent sampling in many drainages, especially inremote areas. There can be little doubt, though, that bull trout are muchless abundant in the study area now than formerly, to the extent that thespecies has virtually disappeared from much of its former habitat in theCCRE.

The bull trout is internationally recognized as vulnerable throughout itsrange (Williams et al. 1989), and an application for listing it for protectionunder the Endangered Species Act has been made in the USA. Its status inCanada has been under review by COSEWIC since at least 1988 (Campbell1988:84, 1993:398). Alberta has recently implemented a specialmanagement and recovery plan for bull trout (Berry 1994).

Brook trout Ñ Native to eastern North America, brook trout have beenintroduced into every major drainage of the CCRE (Figure 8). They havebecome widely established in all drainage basins in the study area, withthe possible exception of the Columbia mainstem and its immediate

28

Figure 7. The present disrtributionof bull trout in the CCRE.

Figure 8. The distribution ofbrook trout in the CCRE. Allpopulations have been introduced.

The Fishes

tributaries, where the only specimen record I encountered was for SuccourCreek (Fielden et al. 1992). Brook trout constitute a threat to weakened bulltrout populations where the two occur together by producing hybridswith that species that rarely reproduce (Leary et al. 1983, 1993).

Dolly Varden Ñ True Dolly Varden charr (as opposed to misidentifiedbull trout) occur within the CCRE in just one location: Chester Lake, in theextreme headwaters of the Kananaskis drainage1, Bow River basin.According to Nelson and Paetz (1992), who discussed the origin of thisstock at length, these fish were introduced in 1974, and are the northernform of the species.

Dolly Varden and bull trout can hybridize and produce viable offspring(Cavender 1978, Haas and McPhail 1991). Chester Lake has a permanentoutlet that could easily allow individuals of this stock to escapedownstream to Smuts Creek via Mud Lake reservoir, and from thereultimately to the rest of the Spray drainage and to the Bow River. Therealready are in fact credible angler reports of northern Dolly Varden in theupper Smuts Creek drainage (Jim Stelfox, Alberta Fish and Wildlife,personal communication 1994). A failure of the Mud dam, diversions bythe utility company or angler releases would allow access to Smith-Dorrien Creek. Once there, Dolly Varden would pose a threat to thegenetic integrity of the trophy native bull trout stock presently beingrestored in Smith-Dorrien Creek. In the worst case, this exotic specieseventually could genetically contaminate all remaining native bull troutstocks in the Bow River watershed not presently protected above barriersto dispersal. These include essentially all those below Bow Falls (BowRiver at Banff) and the Glenmore dam (Elbow River at Calgary), as well asvirtually all remaining stocks in the Highwood drainage (possiblyexclusive of those in the upper Highwood River above a barrier below theCCRE boundary).

Lake trout Ñ Lake trout are believed to be native to a small number oflakes and associated rivers on the east slopes within the CCRE, in theNorth Saskatchewan (Glacier, Outram, Clearwater, Swan lakes, NorthSaskatchewan and Alexandra rivers) and Bow (Minnewanka) drainages(Ward 1974, Nelson and Paetz 1992, Donald and Alger 1993). The species1 This is the natural drainage. A dam on Mud Lake at the height of land now causes

Chester Creek to drain to Smuts Creek, Spray River drainage, Bow basin.

31

The Fishes

has been introduced into Bow and Hector lakes, and Spray Reservoir, allin the Bow drainage; and into the Kicking Horse drainage in Wapta andSherbrooke lakes (Donald and Alger 1993). A record of a single lake troutfrom Moraine Lake (Mayhood and Anderson 1976, Mayhood 1978) was infact a splake (Mudry and Anderson 1977:26): there is no record of laketrout being stocked there (Ward 1974). Non-native lake trout stocks havebeen introduced into waters with native lake trout stocks in LakeMinnewanka (BNP stocking records) and possibly in Swan Lake (SteveHerman, Alberta Fish and Wildlife, personal communication). It ispresently not known whether introduced and native lake trout stocks haveintrogressed where they have come into contact, although the potentialclearly exists for it. Native bull trout have been displaced by lake troutintroduced into Bow and Hector lakes, although native stocks of eachspecies continue to coexist in two relatively pristine CCRE lakes (Glacierand Clearwater, Donald and Alger 1993).

Splake Ñ This hybrid of lake trout and brook trout was produced formany years in the Banff hatchery (Stenton 1950, 1952; Solman et al. 1952;Ward 1974). The hybrid is both viable and highly fertile, capable in turn ofproducing viable, fertile offspring in matings with other splake or witheither parental species (Stenton 1950, 1952; Buss and Wright 1956, 1958;Berst et al. 1980). In this sense it may be considered a hybrid species.Spawning and natural recruitment in the wild occurs (Goldberg et al. 1967,Berst et al. 1980:878-880), and if continued ultimately would produce anintrogressed hybrid swarm in which it would be visually impossible todistinguish certain hybrids from the parental species (cf. Berst et al.1980:843).

Splake were stocked into several CCRE lakes in BNP (Agnes, First Block,Second Block, Boulder, Hector, Herbert, Louise, Minnewanka, Moraine,Mosquito, Turquoise and Two Jack lakes, Bow drainage; Sunwapta Lake,North Saskatchewan drainage), YNP (Duchesnay, Summit and Waptalakes) and KNP (Kootenay Pond) (Ward 1974). The Agnes Lakepopulation reproduced successfully in 1954 after its introduction in 1951(Goldberg et al. 1967, Ward 1974), and persisted until at least 1975,although it apparently was very small then and in 1976 (Mayhood andAnderson 1976:14). The reproductive success and continued existence ofmost other populations is unknown, although the populations in Hector,Minnewanka and Turquoise lakes are thought to be naturally reproducing

32

The Fishes

(Ward 1974:Appendix C, but cf. Ward 1974:12).

Because many of these populations have access via lake outlets to laketrout, brook trout and other splake populations, it is conceivable that laketrout-brook trout genetic introgression has spread beyond the originalstocking sites. This is particularly true in the Bow drainage, where splake,brook trout and lake trout introductions have been numerous and wherebrook trout are widespread and often abundant. All populations of laketrout and brook trout that are or were accessible to splake populationsshould be considered potentially contaminated with the genes of brooktrout or lake trout, respectively.

Cisco Ñ Ciscoes of Great Lakes origin were successfully introduced intoLake Minnewanka (Bow drainage) in 1916-17 (Ward 1974). A stockintroduced into the Spray reservoir in 1953, though initially successful,now appears to be declining toward extirpation (J. Stelfox, Alberta Fishand Wildlife Service, personal communication; Nelson and Paetz 1992).The locality record of cisco in the headwaters of the North Saskatchewangiven by Scott and Crossman (1973:238) requires confirmation. Althoughthe species is under review by COSEWIC for possible listing asendangered (Campbell 1993:398), this evidently refers only to populationsin lakes Ontario and Erie (cf. Campbell 1992:4). The Great Lakes stock inLake Minnewanka thus might have special value for preservation if it wasderived from either of these lakes.

Lake whitefish Ñ Lake whitefish were introduced unsuccessfully intoLake Minnewanka (Bow drainage) in 1953-58 (Ward 1974:22). Nelson andPaetz (1992:239) suggest, however, that this introduction may have givenrise to a lake whitefish population now existing downstream in Ghostreservoir on the Bow mainstem.

Pygmy whitefish Ñ The Kicking Horse River at Field is the type localityfor the pygmy whitefish (Eigenmann and Eigenmann 1892, Eigenmann1895). The Columbia mainstem drainage within the CCRE harbours it aswell: it has been recorded from Kinbasket Lake and the Blaeberry River(McCart 1970). The pygmy whitefish has not been reported from any ofthe other major drainages of the CCRE. The type locality at Field has beenrepeatedly abused by construction activities and poor sewage disposalpractices (M. Pole, personal communication 1991). The impact of theseabuses on the the originally-described population is not known.

33

The Fishes

Mountain whitefish Ñ Like bull trout, mountain whitefish arewidespread in all major drainages in the CCRE (Figure 9). Most stocksappear to be intact. Mountain whitefish do seem to have disappeared fromJumpingpound Creek within the study area sometime between the mid-1960s and the late 1970s, however (Cunningham 1961, 1962, 1964; cf. Trippet al. 1979, Nibourg 1985). In addition the Bow Lake population may havedeclined markedly since the 1930s, judging from catch-per-unit-effortcomparisons (Donald 1987:551, Donald and Alger 1993:242).

Arctic grayling Ñ Arctic grayling were recently introduced into theCCRE: in Big Iron Lake and Bear Pond, close together in the extremeheadwaters of the Willow Creek drainage, Oldman River system. The BigIron population is reproducing, and is expected to maintain itself (J.Stelfox, Alberta Fish and Wildlife Service, personal communication).

Trout-perch Ñ This species has been recorded near or in the CCRE inJumpingpound Creek, and near the study area in Threepoint, Fish andFisher creeks, Bow drainage; and near or in the CCRE in the James Riverdrainage, Red Deer system (Henderson and Peter 1969, MonencoConsultants Limited 1982). The species might occur within the study areain these and other streams on the east slopes (Nelson and Paetz 1992:299).

Burbot Ñ Burbot are known to occur within the CCRE in the Bow, RedDeer and North Saskatchewan drainages (Henderson and Peter 1969,Nelson and Paetz 1992, Alberta Fish and Wildlife Service file data), and inthe mainstem Columbia (Eigenmann 1895, Carl et al. 1959:142). Thespecies does not appear to occupy the upper Kootenay drainage (Scott andCrossman 1973, Lee et al. 1980, Alger and Donald 1984), nor are thererecords for the Kicking Horse drainage above Wapta Falls (M. J. Pole,personal communication).

Western mosquitofish Ñ This species was introduced in 1924 into theCave and Basin hotsprings to control mosquitoes, and now is thrivingthere (McAllister 1969, Nelson 1983). It has been cited as one of severalexotic fishes possibly contributing to the extinction of the Banff longnosedace endemic to this site (Renaud and McAllister 1988). It may not be ableto successfully reproduce in the cold waters of the Bow drainage beyondthe immediate hotsprings area, although it has been reported under the ice

34

Figure 9. The native distribution ofmountain whitefish in the CCRE.

The Fishes

of the Cave and Basin marsh in winter (Nelson and Paetz 1992).Mosquitofish once were found in the warm spring across the Bow River atThird Vermilion Lake (Ward 1974), but this population has not been seenfor some time (Sentar 1992). The mosquitofish is found nowhere else in theCCRE.

Sailfin molly Ñ This tropical species was introduced into the Cave andBasin hotsprings marsh at Banff in 1960 by a local aquarist (Nelson andPaetz 1992). It has since established a self-sustaining population there, butprobably cannot survive beyond the immediate vicinity of the hotspringwaters. The sailfin molly has been cited as one of several exotic fishespossibly contributing to the extinction of the Banff longnose dace endemicto this site (Renaud and McAllister 1988).

Guppy Ñ Guppies introduced by local aquarists into the marsh at theCave and Basin hotspring and the warm spring at Third Vermilion Lakehave not survived (Ward 1974:31, Nelson and Paetz 1992:382, Sentar 1992).

Green swordtail Ñ This tropical fish initially was successfully introducedby aquarists into the marsh at the Cave and Basin Hotsprings, but hassubsequently disappeared (Ward 1974, Nelson and Paetz 1992).

Brook stickleback Ñ Brook sticklebacks occupy the Bow, Red Deer andNorth Saskatchewan watersheds in the CCRE (Henderson and Peter 1969,Nelson and Paetz 1992, L. Rhude, personal communication). The speciesdoes not occur west of the Continental Divide in our area (Carl et al. 1959,Lee et al. 1980).

Mottled sculpin Ñ Fielden et al. (1992) reported finding this species inone or more unspecified tributaries to Kinbasket Lake. Some of their studystreams were within the CCRE. The record needs confirmation. This is ataxonomically difficult and variable species in need of revision, beingreadily confused with the shorthead sculpin, Cottus confusus (Peden et al.1989), recognized by COSEWIC as a threatened species (Campbell 1993). Ifthe identification is correct, it marks a major range extension of more than300 km above the closest previously-identified population in the Columbiasystem (Peden et al. 1989).

36

The Fishes

Slimy sculpin Ñ This widespread sculpin is absent from east slopestreams within the CCRE (Nelson and Paetz 1992). It is known west of thedivide from the mainstem Columbia, Kootenay and Kicking Horsedrainages, including the latter drainage above Wapta Falls (Eigenmannand Eigenmann 1892, Eigenmann 1895, Carl et al. 1959, Ward 1974, Algerand Donald 1984).

Torrent sculpin Ñ The torrent sculpin has been reported from justoutside the CCRE in the Columbia mainstem at Kinbasket Lake, and in theKootenay drainage from the Canal Flats area (Lee et al. 1980). The onlyrecords for the species within the CCRE, however, are in Emerald Lakeand Ice River, above Wapta Falls in the Kicking Horse drainage (M. J. Pole,personal communication 1991; Pole 1991 cited in Triton EnvironmentalConsultants Ltd. 1992). Torrent sculpins are not known to occupy streamseast of the Continental Divide anywhere within their range (Lee et al.1980:822).

Spoonhead sculpin Ñ There are records of this species in the study area inthe North Saskatchewan and Red Deer drainages, but the nearest recordsfor the Bow drainage are downstream from Cochrane and in lowerThreepoint Creek (Henderson and Peter 1969, Ward 1974, Nelson andPaetz 1992). The spoonhead sculpin has not been found anywhere in thePacific drainage (Lee et al. 1980).

Sunfish (unidentified) Ñ RL & L Environmental Services Ltd. (1993)reported collecting young centrarchids of the genus Lepomis in ColumbiaLake, on the mainstem Columbia River. This population clearly hasresulted from an introduction.

Smallmouth bass Ñ Ward (1974:2), citing a Òmemo on old National Parkfiles,Ó reported the introduction of this species to Lake Minnewanka in1901-02. This is undoubtedly the failed introduction to this lake of Òsome500 parent Black Bass and some 3,000 fingerlingsÓ referred to by theDominion Alberta and Saskatchewan Fisheries Commission of 1910-11Òsome ten years agoÓ (Prince 1912:48). The term Òblack bassÓ wascommonly used for both this and the following species (Prince et al.1912:48-49).

37

The Fishes

Largemouth bass Ñ Windermere Lake, on the Columbia River mainstem,is said to hold this introduced species (W. Westover, personalcommunication). Prince et al. (1912:38) did not believe that introducingbass to this lake would be successful, but raised a concern that if successfulthe species might spread elsewhere in the system Òwhere they were notdesired.Ó

Convict cichlid Ñ the convict cichlid, a tropical species, was introducedwithout success into the marsh at Cave and Basin Hotsprings, BNP (Bowdrainage) by local aquarium hobbyists (Nelson and Paetz 1992:385).

African jewelfish Ñ This is another of the many tropical species that havebeen introduced into Cave and Basin hotsprings in BNP (Nelson and Paetz1992). The African jewelfish probably is not capable of surviving outsidethe confines of the marsh fed by the hotsprings.

Freshwater angelfish Ñ Nelson and Paetz (1992:385) state that thistropical species, once introduced into the marsh at the Cave and BasinHotsprings, BNP (Bow drainage) has not been seen since 1968, andprobably did not survive for long.

38

Discussion

Discussion

Faunal Loss: Causes and Consequences

Native fish stocks have not fared well in the CCRE. Of the 33 speciesbelieved to be native to the study area or adjacent connected waters, atleast 10 have suffered impacts of significant conservation concern (Table3). Only four native species could be considered to have benefitted overallfrom human intervention. It is worth elaborating on some of the moresignificant concerns, emphasizing two of the most prominent species.

Westslope Cutthroat Trout

Westslope cutthroats like other interior subspecies have been decimatedthroughout their range by the cumulative effects of overexploitation,habitat damage, displacement by introduced species and geneticintrogression from introduced black-spotted trout stocks (e.g., Nelson1965; Reinitz 1977; Bjornn and Liknes 1986; Allendorf and Leary 1988;Liknes and Graham 1988; Trotter 1991; Behnke 1992). Of these,introgressive hybridization with introduced rainbow trout and non-nativecutthroat stocks is often considered to have been the factor mostresponsible for the destruction of this subspecies.

An exotic subspecies, the Yellowstone cutthroat, has been widelyintroduced in the national parks of the CCRE, exposing many nativewestslope cutthroat populations to introgression (McAllister et al. 1981). Agenetically-pure population of Yellowstone cutthroats still exists in TaylorLake (Bow drainage, BNP, McAllister et al. 1981), and probably also inseveral other park lakes. Cutthroats from the state of Washington, possiblycoastal cutthroats, also were stocked in the Bow drainage for several years(Techman Ltd. 1980).

But I believe by far the most serious hybridization problem arose from theintroduction of rainbow trout, widely, persistently and in large numbers,into waters that had been depleted of their native cutthroat stocks, withthe results described in this report. Complaints of widespread overfishingand wanton destruction of trout populations by almost every conceivable

39

Discussion

Table 3. Summary of the status of native fishes in the CCRE. Designations refer to the study areaonly unless stated otherwise. Only species of some conservation concern are included.

Species Status

Pacific lamprey probably extirpated, but conceivably a landlocked stock still exists

white sturgeon status unknown: no longer anadromous, stock may be extirpated

chiselmouth rare at best, COSEWIC review re possible status as vulnerable in BC

lake chub range expanded through introductions, commonly hybridizes withlongnose dace in reservoirs

longnose dace range expanded through introductions, commonly hybridizes withlake chub in reservoirs

Banff longnose dace extinct

leopard dace possibly not in study area; uncommon, vulnerable to effects of dams

longnose sucker several suspected introductions into non-native waters; commonlyhybridizes with white sucker in reservoirs

white sucker some suspected introductions into non-native waters; commonlyhybridizes with longnose sucker in reservoirs

largescale sucker some suspected introductions into non-native waters

westslope cutthroat trout most native stocks destroyed, all remaining native stocks in nativehabitats endangered

rainbow trout any possible anadromous stocks destroyed; native status unknown &threatened by introgression from non-native stocks

sockeye salmon any native anadromous stocks extirpated

chinook salmon extirpated

bull trout extirpated from some waters, severely depleted throughout area;threat of introgression from introduced Arctic charr & Dolly Varden?

lake trout native stocks rare, 2 possibly introgressed with introduced stocks

pygmy whitefish population of type locality threatened by continued abuse of habitat

means in the CCRE were voiced shortly after the arrival of the CPR in themid-1880s, and prompted immediate demands for hatchery restocking(Whitcher 1887; McIllrie and White-Fraser 1983, re 1890; Prince et al. 1912).The hatcheries were built and trout both native and exotic, especiallyrainbow trout, were transplanted or introduced throughout the CCRE(Miller and Macdonald 1950, Ward 1974, Donald 1987, Mayhood 1992a:9-14). Obvious rainbow-cutthroat hybrids were widespread by at least 1947

40

Discussion

(Miller and Macdonald 1950, Miller 1950).

As of this writing, only 12 existing cutthroat stocks in the CCRE have beendemonstrated by reliable biochemical means to be genetically pure. Theseare in Upper Block, Elk, First Fish, Second Fish, Third Fish, Marvel,Mystic, Taylor, Lower Twin (all in BNP) and Picklejar #4 lakes (Bowdrainage); Floe (KNP) and Connor lakes (Kootenay drainage) (McAllisteret al. 1981, Leary et al. 1985, Carl and Stelfox 1989). The Marvel andConnor lakes stocks have been widely transplanted in Alberta and BC,respectively. While many of the populations arising from these transplantsmay remain genetically uncontaminated, they represent only two geneticlines, each of which is remarkably homogeneous (i.e., highly homozygous)(McAllister et al. 1981, Carl and Stelfox 1989).

At least four of these pure stocks certainly are not indigenous to the watersin which they are now found. As noted above, the Taylor Lake populationis an exotic (non-Canadian) subspecies almost certainly derived fromYellowstone Lake, Wyoming (McAllister et al. 1981). The Marvel Lakepopulation was transplanted from the nearby Spray Lakes (Rawson 1947,Techman Ltd. 1980). The Floe Lake population was not native to that lake,and almost certainly no longer exists due to lack of natural recruitment(Donald and Alger 1984). Floe Lake was last stocked with cutthroats froman unknown source in 1968 (BNP stocking records cited by Techman Ltd.1980). The Upper Block Lake population is self-maintaining, but is knownto have been stocked in 1957 (source unknown) and 1958 (Cranbrookhatchery) (Ward 1974, Techman Ltd. 1980). This lake lies over 300 metresabove the valley floor at the top of a cliff, lacks a surface outlet, andcertainly was inaccessible to colonizing native fish.

In addition to the above, there are reasonable concerns about whethersome of the other genetically-pure populations are truly indigenous to thelakes in which they are found. For example, Lower Twin Lake does have apermanent surface outlet that would appear to be passable to fish, and aself-maintaining cutthroat population. This lake, however, was eitherstocked directly with cutthroats from an unrecorded source in 1953 (Ward1974), or was accessible to cutthroats from an unrecorded source stockedin the upper Twin Lake that same year (Techman Ltd. 1980). Becauseunrecorded introductions were commonly made in lakes of the nationalparks for many years, at this time we cannot be entirely confident that any

41

Discussion

of the known genetically-pure populations are actually native to thewaters in which they are now found.

All or nearly all of the native stream populations in the Bow, Kootenayand upper Kicking Horse drainages have been exposed to introgressionfrom introduced rainbow trout (Figure 2, cf. Figure 4), and sometimesfrom introduced cutthroat stocks. Most remnant populations throughoutthe mainstem Bow, Jumpingpound, Elbow, Sheep and Highwood basinsshow the effects of introgression with rainbow trout in their colouring,spotting patterns and morphology. Difficulties in distinguishing betweenintroduced rainbow and native cutthroat trout in these drainages, evidentat least by 1947-1949 (Miller and Macdonald 1950:41-44, Miller 1950), werealmost certainly due to widespread introgressive hybridization.

The existing lake populations in the Bow and other CCRE drainages underAlberta jurisdiction are mostly introduced populations derived from asingle, genetically-depauperate Spray Lakes stock via Marvel and Joblakes (Rawson 1947, Techman 1980, McAllister et al. 1981, Carl and Stelfox1989). Introduced populations in the North Saskatchewan and Red Deerdrainages have greatly expanded the range of westslope cutthroat trout inthe CCRE. Isolation of transplanted westslope cutthroats above major fallsin the Ram, Palliser and Fording river drainages may serve to protect atleast two or three artificial stream stocks from genetic contamination infuture, if in fact these populations originated from geneticallyuncontaminated stocks in the first place.

The westslope cutthroat trout is so genetically distinct from all othercutthroats that it should be treated as a separate species in its own right(Leary et al. 1987, Allendorf and Leary 1988). Given the widespreaddestruction of its stocks throughout its range, it is critical to preserve allremaining genetically-pure stocks of this fish.

A high proportion of the total genetic diversity of westslope cutthroattrout is represented by unique alleles existing at high frequency inisolated, genetically-uniform populations (Leary et al. 1987). This isparticularly true of the remnant stocks in the Bow basin, which areespecially divergent genetically among themselves and as a group fromstocks elsewhere throughout the range of the subspecies (Leary et al.1985). For this reason, preserving the remaining genetic integrity of thissubspecies requires preserving as many local native populations as

42

Discussion

possible (Leary et al. 1987, Allendorf and Leary 1988), those in the Bowdrainage being especially important to protect in this respect.

Bull Trout

Almost everything about the biology of the bull trout makes it vulnerableto extirpation, so the possible causes for the decline of this species in theCCRE are numerous (Mayhood 1992b:186-205, Rieman and McIntyre1993). Bull trout mature late, so become vulnerable to fishing mortalitywell before the age of first spawning. They seem to favour groundwaterdischarge areas, which usually have a limited distribution in mostwatersheds, as spawning and rearing habitat. Incubating bull troutembryos, early fry and juveniles are highly sensitive to silting and otherbottom disturbances. Because many stocks home strongly to their nativestreams, a lost stock is not soon replaced. The adults are large (attractingangler interest), strongly predaceous and easily caught by angling. Theyare highly visible and easily poached in their tiny spawning streams.Adults undergo long spawning migrations in many stocks, and aretherefore vulnerable to being isolated by dams from their spawningstreams.

Depletion or complete loss of certain stocks in the Clearwater River appearto have been due to overexploitation rather than to habitat damage ordisplacement by exotics such as brown trout (Allan 1980). In contrast thenearly complete eradication of bull trout from the Kananaskis Riversystem below Lower Kananaskis reservoir evidently had severalcontributory causes, including displacement by brown trout (whichapparently were favoured in the widely fluctuating water levels of thishydroelectric dam-dedicated system) and overexploitation (Nelson 1965).It also seems likely that certain of the dams blocked spawning runs.Elsewhere depleted bull trout stocks have been linked to habitatdisruption (especially siltation and increased bedload movement fromdisruptions in the watershed) and displacement by brook trout, whichoverwhelm bull trout populations by hybridizing with them to createoffspring which fail to reproduce (Rieman and McIntyre 1993, Leary et al.1993).

43

Discussion

It is sometimes held or implied that overexploitation is the chief factorresponsible for the decline of bull trout in Alberta (e.g., Roberts 1987, 1991,1993; Palmer 1994). Clearly other factors are important in many cases. Butmore to the point, looking for a chief factor for population declines ismisleading for future conservation efforts, because factors interact. Forexample, restoring a trout stock depleted by overfishing will be impossibleif its critical habitat is not protected, or if introduced competitor stocks,now with the upper hand, occupy the habitat. Blocking access to spawningsites with a poor culvert or a dam will destroy the population whether it isfished or not. In managing, protecting or restoring fish stocks care must betaken to manage with the whole ecosystem in mind.

The losses and depletion of so many bull trout stocks in the CCRE is aserious threat to the continued existence of the species in the study area.Like westslope cutthroat trout, bull trout where they have been adequatelystudied show little within-population genetic variation, but significantgenetic differences among populations (Leary et al. 1993). If the same istrue of stocks in the CCRE, preserving the genetic diversity of bull trouthere will require the continued existence of populations throughout theregion.

Other Species

The causes of damage to the remaining native species adversely impactedby human activities in the CCRE are sometimes clearer to discern. Knownor potential anadromous stocks of Pacific lamprey, white sturgeon,steelhead, sockeye salmon and chinook salmon have been destroyed (orpossibly landlocked in some cases) by barrier dams on the Columbia Rivermainstem. Similarly, dams may now exclude lake sturgeon from enteringEast Slopes waters of the CCRE. Dams and impoundments have createdfavourable conditions for the invasion or introduction of manynonindigenous stocks (e.g., the spawning invasion of the upper KootenayRiver in KNP by thousands of kokanee introduced into Koocanusareservoir) and the extensive hybridization of several species (e.g., Nelson1965, 1966, 1973).

The mechanism causing the extinction of the Banff longnose dace in Caveand Basin Hotspring is an example of how complex the causes of faunal

44

Discussion

loss can be. Evidently introductions of several tropical species, periodiccessation of flows arising from the use of the hotspring as a public bath,periodic sewage spillage and introgressive hybridization of the weakenedhotspring stock with another native subspecies all played a role in thedemise of the subspecies (Renaud and McAllister 1988, Miller et al. 1989).

The New Fauna

In contrast to native fish stocks, many introduced stocks have done well inthe CCRE. Of 21 species introduced to the study area or adjacentconnected waters, 13 have managed to sustain populations somewherewithin it (Table 2). At least eleven are of significant conservation concern(Table 4). None have produced significant new sport fisheries that couldnot have been provided by native or transplanted native stocks.

Table 4. Summary of the status of introduced fishes in the CCRE. Only species of someconservation concern are listed.

Species Status

Yellowstone cutthroat trout introduced widely into Bow drainage westslope cutthroat range, some introgression identified

kokanee widespread, abundant, invades Kootenay National Park in large numbers

brown trout introduced & expanding range in east slopes mainstems

Arctic charr introduction apparently unsuccessful; some stocks possibly introgressed withnative bull trout stocks

brook trout widespread & well-established exotic in all major drainages; competitivethreat to native cutthroat and bull trout

Dolly Varden 1 population, hybridization threat to 1 remnant bull trout population

lake trout introduced stocks displaced bull trout in at least 2 lakes

splake status unknown, introgression threat to some introduced lake & brook trout

cisco 1 introduced stock, potentially valuable for future reintroduction intonative Great Lakes waters where stocks may be endangered

western mosquitofish exotic contaminant of a unique hotspring habitat, possiblecontributor to extinction of Banff longnose dace

sailfin molly exotic contaminant of a unique hotspring habitat, possiblecontributor to extinction of Banff longnose dace

African jewelfish exotic contaminant of a unique hotspring habitat

45

Discussion

The losses of so many native fish stocks, and the successful introductionand transplantation of non-native stocks, has transformed the fish fauna ofthe CCRE. The bull trout and cutthroat trout assemblage characteristic ofthe Bow drainage has been replaced by a trout fauna in which brook troutpredominate above Bow Falls. Below Bow Falls brown and brook trouthave completely replaced the native cutthroat and bull trout. The samenative trout fauna of the lower Bow tributaries of Jumpingpound Creekand the Elbow, Sheep and Highwood rivers is now largely replaced byexotic brook and hybrid rainbow-cutthroat trout. The unique Cave andBasin microecosystem has been transformed into little more than a tropicalfish pond.

Introduced cutthroat trout dominate the formerly fishless Ram Riverdrainage above the barrier falls, and are widespread in the upper NorthSaskatchewan drainage. Brook trout are pervasive almost everywhereexcept the Columbia River mainstem and its direct tributaries, andintroduced rainbow trout occupy large portions of the Kootenay drainage.The Columbia has lost its anadromous stocks of chinook salmon that soimpressed the first European travellers. The native fish fauna of the upperKicking Horse appears to have all but vanished, replaced by brook trout,and introduced rainbow and cutthroat stocks. Introduced brook andcutthroat trout occupy many dozens of formerly fishless mountain lakes.Several sucker and minnow species are expanding their ranges throughillegal introductions. In almost every drainage the abundant supplies oflarge native trout reported by early anglers have been replaced by largereaches of empty stream habitat.

In short, the present fish fauna of the CCRE is overwhelmingly artificial,depauperate and dominated by exotic stocks. The native fauna has beenlargely obliterated; only tiny fragments remain.

Realistic Response:Ecosystems for Fish Conservation & Management

The pervasiveness of the changes in the fish fauna of the CCREsummarized above make it clear that the ecological integrity of the studyarea has already been lost. The question is no longer one of maintaining

46

Discussion

ecological integrity of the region, but of retaining whatever remnants wecan, perhaps restoring to some limited extent some small portion of theCCRE. The situation is analogous to a battle that has been decisively lostwith appalling casualties, and the approach must be one of triage.

triage: 1 the act of sorting according to quality. 2 theassignment of degrees of urgency to decide the order oftreatment of wounds, illnesses, etc. Ñ Concise OxfordDictionary, 8th edition, 1990

Because the damage is so widespread and profound, and because relateddamage to other biota and habitats exists throughout the CCRE, it wouldprove futile to rely solely on the population-based approaches oftraditional fishery biology to address the problems. An ecosystem-basedapproach is needed. But is the current conception of the Central CanadianRockies Ecosystem suitable for dealing with the ecological disintegrationof its fish stocks?

The Continental Divide separates two distinct native fish faunas within theCCRE. Of 33 species that are believed to be native to the CCRE or itsimmediate surroundings, 13 are native exclusively to waters on the westslopes, and 13 are native only to the east slopes. Just seven (lake chub,longnose dace, longnose sucker, westslope cutthroat trout, bull trout,mountain whitefish and burbot) occupy waters on both sides, but three ofthese (lake chub, longnose dace and burbot) are represented by differentmorphotypes or subspecies on either side of the divide (McPhail andLindsey 1970:244, 1986:628-629; Renaud and McAllister 1988:108-110).Thus only 11% of the native fish fauna (four of 36 distinguishable types) iscommon to waters on both sides of the divide in the CCRE.

It is thus obvious that the Central Canadian Rockies Ecosystem aspresently demarcated does not comprise an appropriate unit ecosystem forfish conservation and management. At the very least a useful naturalecosystem for such purposes should include all waters within whichnatural genetic interchange among aquatic populations is possible. Itshould also be functionally integrated such that the physical, chemical andbiological elements and processes with which the fish stocks interact are

47

Discussion

included within their ecosystem boundaries, while those with which thereis little or no interaction are excluded. The fishes of the NorthSaskatchewan drainage, for example, do not interact with those in theKootenay drainage. The physical, chemical and biological elements andprocesses of the Red Deer basin are not an integral part of the ecosystem offishes in the Columbia drainage. What is of direct and intimate relevanceto the Red Deer drainage fish stocks are the physical, chemical andbiological elements and processes of the Red Deer basin. The NorthSaskatchewan basin fishes interact among themselves, and not with thoseon the far side of the Continental Divide.

For these reasons discrete drainage basins are especially useful naturalecosystem units for conserving and managing aquatic organisms and theirhabitats (Hynes 1975, Lotspeich 1980, Stanford and Ward 1992).Watershed ecosystems are inherently organized hierarchically, withrelated basins being nested together within common larger basins, a usefulmanagement arrangement. Several such watershed ecosystemsincorporating the presently-defined CCRE have been used to organize thedata for this report: the North Saskatchewan, Red Deer, Bow, Kootenayand Columbia rivers. The upper Kicking Horse drainage is a naturalecological subunit of the Columbia, with unique yet related features tothose in the larger basin. Likewise the Ram drainage is a unique butintegral part of the larger North Saskatchewan drainage.

Banff, Yoho and Kootenay national parks taken together can be thought ofas the core area supporting the conservation of ecological integrity in thelarger CCRE. Yet they would far better serve the purpose for fishconservation if managers organized their activities around watersheds asthe unit ecosystems. For example the Kootenay drainage basin withinKootenay National Park should be thought of as a protected area of thegreater Kootenay River Ecosystem. Somewhat similarly the upper KickingHorse watershed within Yoho and Kootenay national parks is in fact aprotected area for conserving as a whole this isolated unit ecosystem, aunique part of the Greater Columbia River Ecosystem. In fact, the majorwatersheds within these three national parks are headwater protectedareas for no fewer than six regional ecosystems. In addition to the two justmentioned, these are the regional ecosystems of the North Saskatchewan,Clearwater, Red Deer and Bow river catchments. Similar roles can readilybe perceived for the several provincial protected areas in the region.

48

Discussion

The proposed regional watershed ecosystems and their headwaterprotected areas within the national parks are illustrated in Figure 10.Downstream boundaries are not shown for most, acknowledging the factthat sharp, ecologically-meaningful boundaries do not exist at the outletsof drainage basins except at barrier waterfalls.

The open-ended nature of these ecosystems forces us to view our aquaticconservation problems in a much more realistic context than the presentenclosed, sharp and entirely arbitrary boundaries of the CCRE allow us todo. Instead of thinking of the CCRE as a tidy patch of landscape that wemight eventually be able to control, we see it as vast open area from whicheverything we do within the region unavoidably leaks out. The nationalparks are shown in their true context, not as the centre of the regionalecosystem, but as rather small, marginal, out-of the-way adjuncts to muchlarger unit ecosystems.

The open-ended watershed model of the CCRE makes it clear that theparks can play only a limited role in protecting the regional aquaticecosystems. Certainly the headwater areas within the national parks mustbe managed in such a way that the ecological integrity of the downstreamwatershed is not compromised. But most ecosystem protection must takeplace primarily outside the parks. In such ways as this, the watershedecosystem view of the aquatic ecosystems of the CCRE will help us toconduct triage realistically.

49

Figure 10. Suggested boundaries for the CCRE forfish conservation and management. Lines mark theboundaries of major regional watershed ecosystems.A - Columbia Mainstem, B - Upper Kicking Horse,C - Upper Kootenay, D - North SaskatchewanMainstem E - Upper Ram, F - Clearwater, G - RedDeer, H - Bow

References Cited

References Cited

Alberta Forestry Lands and Wildlife. 1992. Provincial base map 1:1,000,000. Alberta Land InformationServices Division, Edmonton, AB.

Alger, D. J. and D. B. Donald. 1984. Limnological studies in Kootenay National Park. Part 2: the streams.Report prepared for Parks Canada by the Canadian Wildlife Service, Edmonton, AB. 112 p.

Allan, J. H. 1980. Life history notes on the Dolly Varden char (Salvelinus malma) in the upper ClearwaterRiver, Alberta. Manuscript report, Fish and Wildlife Division, Alberta Energy and NaturalResources, Red Deer, AB. 59 p.

Allendorf, F. W. and R. F. Leary. 1988. Conservation and distribution of genetic variation in a polytypicspecies, the cutthroat trout. Conservation Biology 2:170-184.

Ambrosi, J. 1987. Fly fishing the Canadian Rockies. Rocky Mountain Books, Calgary, AB. 218 p.

Anderson, R. S. and D. B. Donald. 1978. Limnological survey of some small lakes in the vicinity of theCascade trail, Banff National Park. Report prepared for Parks Canada by the Canadian WildlifeService, Calgary, AB. 162 p.

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